传粉动物多样性的保护与农业景观传粉服务的提升

传粉动物为许多植物尤其是作物提供了重要的传粉服务, 在保障全球粮食安全和人类福祉、缓冲气候变化对作物产量的影响等方面都发挥着重要的作用。然而来自全球土地利用变化、化学农药使用、外来物种入侵及气候变化等的威胁, 导致传粉动物的多样性下降并造成了依赖动物传粉的作物产量和品质的下降。针对这一情况, 作者提出了农业景观传粉动物多样性保护和利用的3种主要途径: (1)改善生产管理, 例如减少化学农药的使用、适当地采取有机种植; (2)促进景观多样性, 包括创建适宜野生传粉者的半自然生境、保护高价值的自然生境、作物多样化、合理配置资源和生境的空间分布; (3)加强对本土传粉动物的保护和开发利用。文章最后提出, 为进一步提升传粉服务, 还需加强对传粉者的生物学特征、传粉服务的需求与供给现状、影响传粉动物多样性和传粉服务的农作措施和景观因素等方面的研究。     

Pollinator diversity and specialization in relation to flower diversity

In the face of global decline in biodiversity, the relationship between diversity and species interactions deserves particular attention. If pollinators are strongly dependent on floral diversity due to mutual specialization, declines in plant diversity, e.g. caused by land use intensification, may be associated with linked extinctions of pollinators. However, the general extent of pollinator specialization is still poorly known. To explore the dependence of local bee and hoverfly communities on flower diversity, we recorded flower supply and flower‐visiting insects on 27 meadows with varying flower diversity in southern Germany and analyzed (a) whether the diversity of flower visitors is correlated with flower diversity, (b) whether the degree of dietary specialization of flower visitors changes with flower diversity and (c) whether flower preferences of individual flower visitor species are constant or variable between different communities. Flower–visitor interaction webs were compiled during a single day on each meadow. This approach prevents relating pollinator species to flowers they never encounter because of non‐overlapping phenology or spatial segregation. (a) Flower diversity and flower visitor diversity were positively correlated. (b) Flower visitor assemblies were significantly specialized at a relatively high level, contrasting to the opinion that plant–pollinator webs are highly generalized, and providing a possible explanation for the positive diversity correlation. However, the level of specialization did not change significantly across the gradient of flower diversity, suggesting that pollinators are partitioned to a similar extent in each meadow. (c) In the analysis of ten common flower visitor species previously categorized as generalists, strong evidence was found for both, consistent preferences and preferences that differ between sites. These results indicate a flexibility in flower preferences and a dynamic resource partitioning among pollinators. Generally, our findings highlight the complexity of plant–pollinator interactions and confirm the importance of flower diversity for bee and hoverfly communities.     

Pollinator behavior and deceptive pollination: learning process and floral evolution

Some species of flowering plants engage in nonmodel deceptive pollination, attracting pollinators by large nonmimetic floral displays and providing no reward. Pollinators can learn to avoid deceptive plants and to favor nectariferous species. The reproductive success of these species is expected to be density dependent for two opposite reasons: the commoner cheating flowers are, the easier they are to avoid and the lower the quality of the patch, making it more difficult to recognize that unrewarding flowers are not profitable. When a deceptive species is made up of multiple floral variants, pollinators' learning could decrease the reproductive success of any particularly common floral variant. Within a population of deceptive plants, mean reproductive success could, therefore, vary with the number of floral variants. We investigate these three hypotheses by modeling the behavior of pollinators foraging in communities of deceptive and rewarding flowers. Simulations revealed that the reproductive success of deceptive flowers varies in a density-dependent manner and that floral variants can be submitted to negative frequency-dependent selection. We compare density dependence in nonmodel deceptive species to what is expected in Batesian mimics and discuss possible selection of morphological variants. Finally, we survey how pollinators' learning capacities can make mean reproductive success depend on morphological variability within a population.     

Effects of grazing intensity on pollinator abundance and diversity,and on pollination services

1. Pollinating insects provide important ecosystem services and are influenced by the intensity of grazing. Based on the Intermediate Disturbance Hypothesis (IDH), pollinator diversity is expected to peak at intermediate grazing intensities. However, this hump‐shaped relationship is rarely found. 2. The effect of grazing intensity was tested on flower cover, on the abundance and richness of bees, hoverflies and bee flies, and on pollination services to early‐flowering bee‐pollinated Asphodelus ramosus L. For that, we used data on 11 plant–pollinator phryganic communities from Lesvos Island (Greece) widely differing in grazing intensities. 3. Flower abundance and richness showed hump‐shaped relationships with grazing intensity. Grazing affected the abundance and richness of bees and hoverflies directly and also indirectly, through changes in the flower community. Grazing influenced directly the richness but not the abundance of bee flies. Overall, pollinator abundance and richness showed hump‐shaped relationships with grazing intensity, but variations in strength (hoverfly abundance) and direction (bee community) of the effect appeared along the season. Early in the season, grazing increased bee abundance but decreased richness, resulting in increased pollen limitation in A. ramosus. 4. The effects of grazing on pollinators vary with the intensity of the disturbance, generally supporting the IDH, and the timing of land‐use activities may influence pollination services. Management strategies should include moderate grazing levels to preserve overall diversity in this area, however, the conservation of particular early bee or bee‐pollinated species may benefit from reduced grazing in early spring.     

Pollinator population size and pollination ecosystem service responses to enhancing floral and nesting resources

Modeling pollination ecosystem services requires a spatially explicit, process‐based approach because they depend on both the behavioral responses of pollinators to the amount and spatial arrangement of habitat and on the within‐ and between‐season dynamics of pollinator populations in response to land use. We describe a novel pollinator model predicting flower visitation rates by wild central‐place foragers (e.g., nesting bees) in spatially explicit landscapes. The model goes beyond existing approaches by: (1) integrating preferential use of more rewarding floral and nesting resources; (2) considering population growth over time; (3) allowing different dispersal distances for workers and reproductives; (4) providing visitation rates for use in crop pollination models. We use the model to estimate the effect of establishing grassy field margins offering nesting resources and a low quantity of flower resources, and/or late‐flowering flower strips offering no nesting resources but abundant flowers, on bumble bee populations and visitation rates to flowers in landscapes that differ in amounts of linear seminatural habitats and early mass‐flowering crops. Flower strips were three times more effective in increasing pollinator populations and visitation rates than field margins, and this effect increased over time. Late‐blooming flower strips increased early‐season visitation rates, but decreased visitation rates in other late‐season flowers. Increases in population size over time in response to flower strips and amounts of linear seminatural habitats reduced this apparent competition for pollinators. Our spatially explicit, process‐based model generates emergent patterns reflecting empirical observations, such that adding flower resources may have contrasting short‐ and long‐term effects due to apparent competition for pollinators and pollinator population size increase. It allows exploring these effects and comparing effect sizes in ways not possible with other existing models. Future applications include species comparisons, analysis of the sensitivity of predictions to life‐history traits, as well as large‐scale management intervention and policy assessment.     

Synergistic interactions of ecosystem services: florivorous pest control boosts crop yield increase through insect pollination

Insect pollination and pest control are pivotal functions sustaining global food production. However, they have mostly been studied in isolation and how they interactively shape crop yield remains largely unexplored. Using controlled field experiments, we found strong synergistic effects of insect pollination and simulated pest control on yield quantity and quality. Their joint effect increased yield by 23%, with synergistic effects contributing 10%, while their single contributions were 7% and 6%, respectively. The potential economic benefit for a farmer from the synergistic effects (12%) was 1.8 times greater than their individual contributions (7% each). We show that the principal underlying mechanism was a pronounced pest-induced reduction in flower lifetime, resulting in a strong reduction in the number of pollinator visits a flower receives during its lifetime. Our findings highlight the importance of non-additive interactions among ecosystem services (ES) when valuating, mapping or predicting them and reveal fundamental implications for ecosystem management and policy aimed at maximizing ES for sustainable agriculture.     

Forests are critically important to global pollinator diversity and enhance pollination in adjacent crops

Although the importance of natural habitats to pollinator diversity is widely recognized, the value of forests to pollinating insects has been largely overlooked in many parts of the world. In this review, we (i) establish the importance of forests to global pollinator diversity, (ii) explore the relationship between forest cover and pollinator diversity in mixed-use landscapes, and (iii) highlight the contributions of forest-associated pollinators to pollination in adjacent crops. The literature shows unambiguously that native forests support a large number of forest-dependent species and are thus critically important to global pollinator diversity. Many pollinator taxa require or benefit greatly from resources that are restricted to forests, such as floral resources provided by forest plants (including wind-pollinated trees), dead wood for nesting, tree resins, and various non-floral sugar sources (e.g. honeydew). Although landscape-scale studies generally support the conclusion that forests enhance pollinator diversity, findings are often complicated by spatial scale, focal taxa, landscape context, temporal context, forest type, disturbance history, and external stressors. While some forest loss can be beneficial to pollinators by enhancing habitat complementarity, too much can result in the near-elimination of forest-associated species. There is strong evidence from studies of multiple crop types that forest cover can substantially increase yields in adjacent habitats, at least within the foraging ranges of the pollinators involved. The literature also suggests that forests may have enhanced importance to pollinators in the future given their role in mitigating the negative effects of pesticides and climate change. Many questions remain about the amount and configuration of forest cover required to promote the diversity of forest-associated pollinators and their services within forests and in neighbouring habitats. However, it is clear from the current body of knowledge that any effort to preserve native woody habitats, including the protection of individual trees, will benefit pollinating insects and help maintain the critical services they provide.     

Pollinator rarity as a threat to a plant with a specialized pollination system

An increasing diversity of highly specialized pollination systems are being discovered, many of which are likely to be vulnerable to anthropogenic landscape modification. Here, we investigate if a specialized pollination system limits the persistence of Caladenia huegelii (Orchidaceae), an endangered species pollinated by sexual deception of thynnine wasps. Once locally common in part of its geographical range, C. huegelii is now largely restricted to small habitat remnants in urban areas. Pollinator surveys coupled with DNA barcoding detected a single pollinator taxon, a small form of Macrothynnus insignis. Phylogenetic analysis revealed that small M. insignis from within the range of C. huegelii are strongly divergent from other wasp populations, suggesting that some reproductive isolation may exist. Although common in intact landscapes outside the range of C. huegelli, small M. insignis individuals were recorded at only 4% of sites in suitable C. huegelii habitat. Accordingly, reproductive success in C. huegelii was low compared with related Caladenia spp., with 33–60% of populations failing to set fruit in any given year. As such, populations are likely to now persist primarily through individual plant longevity rather than reproduction. Due to the low reproductive success of C. huegelii, ongoing human intervention will almost certainly be needed to sustain the species. Future research will need to focus on optimizing hand pollination to maintain reproduction and high seed fitness. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 511–525.     

天南星科植物的花多样性与传粉策略

天南星科植物具有特殊的佛焰苞花序及多样化的传粉策略, 是研究被子植物花的分化与动植物之间进化生态学联系的理想材料。本文简述了天南星科不同类型的花序结构及其传粉适应意义, 总结了天南星科传粉策略的基本类型与演化历史。天南星科的苞片结构主要包括原始型、外展平面型、直立宽佛焰苞型和直立狭佛焰苞4种类型, 呈现出从简单的片状与外展平面状结构向复杂的立体包裹状的佛焰苞结构演化的趋势。肉穗花序可分为两性花花序、单性花雌雄同序和单性花雌雄异序3种类型, 演化路线为两性花花序→单性花雌雄同序→单性花雌雄异序。天南星科的传粉者主要有鞘翅目、双翅目、膜翅目昆虫, 表现出5种主要传粉策略: 食物报酬型互利传粉、气味吸引型欺骗性传粉、交配场所型互利传粉、产卵场所型互利传粉和致死陷阱型欺骗性传粉。天南星科植物通过花序的形状、颜色、产热以及花部挥发物来吸引传粉者, 其中最主要的挥发物有二甲基硫化物、甲基吲哚化合物、萜类和苯类化合物, 模拟食物或产卵场所信号吸引鞘翅目甲虫和双翅目昆虫为其传粉。天南星科植物的佛焰苞被认为是促进该科物种分化的一个重要结构, 但该性状的演化历史及其与传粉系统分化之间的内在联系尚不明确。利用现代分子生物学技术以及模型模拟等手段, 结合生理生态学方法深入探究传粉事件与天南星科植物的花多样性以及物种分化之间的联系, 有望提升关于植物-传粉者互作与植物的花多样性分化之间关系的认识, 并丰富对被子植物多样性演化相关研究的理解。     

Modelling pollination services across agricultural landscapes

Background and Aims

Crop pollination by bees and other animals is an essential ecosystem service. Ensuring the maintenance of the service requires a full understanding of the contributions of landscape elements to pollinator populations and crop pollination. Here, the first quantitative model that predicts pollinator abundance on a landscape is described and tested.

Methods

Using information on pollinator nesting resources, floral resources and foraging distances, the model predicts the relative abundance of pollinators within nesting habitats. From these nesting areas, it then predicts relative abundances of pollinators on the farms requiring pollination services. Model outputs are compared with data from coffee in Costa Rica, watermelon and sunflower in California and watermelon in New Jersey–Pennsylvania (NJPA).

Key Results

Results from Costa Rica and California, comparing field estimates of pollinator abundance, richness or services with model estimates, are encouraging, explaining up to 80 % of variance among farms. However, the model did not predict observed pollinator abundances on NJPA, so continued model improvement and testing are necessary. The inability of the model to predict pollinator abundances in the NJPA landscape may be due to not accounting for fine-scale floral and nesting resources within the landscapes surrounding farms, rather than the logic of our model.

Conclusions

The importance of fine-scale resources for pollinator service delivery was supported by sensitivity analyses indicating that the model''s predictions depend largely on estimates of nesting and floral resources within crops. Despite the need for more research at the finer-scale, the approach fills an important gap by providing quantitative and mechanistic model from which to evaluate policy decisions and develop land-use plans that promote pollination conservation and service delivery.Key words: Agriculture, bees, ecosystem services, landscape ecology, model, land use, pollinators     

The effectiveness of flower strips and hedgerows on pest control,pollination services and crop yield: a quantitative synthesis

Floral plantings are promoted to foster ecological intensification of agriculture through provisioning of ecosystem services. However, a comprehensive assessment of the effectiveness of different floral plantings, their characteristics and consequences for crop yield is lacking. Here we quantified the impacts of flower strips and hedgerows on pest control (18 studies) and pollination services (17 studies) in adjacent crops in North America, Europe and New Zealand. Flower strips, but not hedgerows, enhanced pest control services in adjacent fields by 16% on average. However, effects on crop pollination and yield were more variable. Our synthesis identifies several important drivers of variability in effectiveness of plantings: pollination services declined exponentially with distance from plantings, and perennial and older flower strips with higher flowering plant diversity enhanced pollination more effectively. These findings provide promising pathways to optimise floral plantings to more effectively contribute to ecosystem service delivery and ecological intensification of agriculture in the future.     

Generating and maintaining diversity at the elite level in crop breeding

Most breeding programs develop elite genotypes that are well adapted to the normal range of environmental conditions in the target production region. These elite lines have similar essential alleles for desirable end use characteristics, agronomics, disease resistance, and adaptation in the target region. The genetic makeup of these elite lines is complex. Intermating among the elite lines will often produce new variability through recombination with minimal risk of introducing new undesirable features, and is the source of most new cultivars. Eventually, this variation will be exhausted and new alleles must be introduced into the elite breeding population. Introducing desirable alleles from exotic germplasm may "pollute" the elite gene pool with undesirable alleles. Exotic germplasm may also disrupt essential allele combinations for adaptation, quality, and agronomic performance. New desirable alleles from exotic germplasm can be introgressed into an elite population in a systematic way through limited backcrossing with a minimal disturbance to the finely tuned elite background. Combining recurrent selection within elite germplasm with a systematic introgression from exotic germplasm in the recurrent introgressive population enrichment (RIPE) system has created an open-ended, continually improving, and sustainable elite population breeding system, which is simple, effective, and a regular source of new cultivars.     

Evolutionarily inspired solutions to the crop pollination crisis

The global decline in insect diversity threatens pollination services, potentially impacting crop production and food security. Here, we argue that this looming pollination crisis is generally approached from an ecological standpoint, and that consideration of evolutionary principles offers a novel perspective. First, we outline that wild plant species have overcome ‘pollination crises’ throughout evolutionary history, and show how associated principles can be applied to crop pollination. We then highlight technological advances that can be used to adapt crop flowers for optimal pollination by local wild pollinators, especially by increasing generalization in pollination systems. Thus, synergies among fundamental evolutionary research, genetic engineering, and agro-ecological science provide a promising template for addressing a potential pollination crisis, complementing much-needed strategies focused on pollinator conservation.     

Pollinator sharing between mass-flowering oilseed rape and co-flowering wild plants: implications for wild plant pollination

Pollinating insects are not only important in wild plant pollination, but also in the production of a large number of crops. Oilseed rape production is increasing globally due to demands for biofuels which may have impacts on pollinating insects which visit the crop and on the pollination services delivered to co-flowering wild plants. In this study, we tested (1) the degree of pollinator sharing between oilseed rape and native wild plants in field margins and hedgerows and (2) the effects of oilseed rape on the quality of pollination service delivered to these wild plants. We found large overlap between flower visitors of wild plants and oilseed rape, but the composition of species overlap differed with respect to each wild plant species. Nearly all individual visitors caught on both the crop and foraging on wild species carried crop pollen, but more than half the insects also carried pollen from wild plants. However, very little oilseed rape pollen was deposited on wild plant stigmas. This shows that (1) oilseed rape overlaps in pollinator niche with most co-flowering wild plants, and (2) crop pollen deposition on wild plant stigmas is low which may indicate that it is unlikely to cause reductions in seed set of wild plants, although this was not measured here. Furthermore, wild plants in field margins and hedgerows are important sources of alternative forage for pollinating insects even when a crop is mass flowering, and we suggest maintenance and augmentation of field margins and hedgerows to provide alternative forage for pollinator conservation to continue provision of pollination services to both crops and wild plants.     

A heterogeneous landscape does not guarantee high crop pollination

The expansion of pollinator-dependent crops, especially in the developing world, together with reports of worldwide pollinator declines, raises concern of possible yield gaps. Farmers directly reliant on pollination services for food supply often live in regions where our knowledge of pollination services is poor. In a manipulative experiment replicated at 23 sites across an Ethiopian agricultural landscape, we found poor pollination services and severe pollen limitation in a common oil crop. With supplementary pollination, the yield increased on average by 91%. Despite the heterogeneous agricultural matrix, we found a low bee abundance, which may explain poor pollination services. The variation in pollen limitation was unrelated to surrounding forest cover, local bee richness and bee abundance. While practices that commonly increase pollinators (restricted pesticide use, flower strips) are an integral part of the landscape, these elements are apparently insufficient. Management to increase pollination services is therefore in need of urgent investigation.     

Pollinator activity and pollination success of Medicago sativa L. in a natural and a managed population

Medicago sativa L. is an important cash crop in the arid region of northwest China. Pollinator activity is an essential aspect of pollination success, but the relationships between pollinator visitation rate and seed set still need further study of M. sativa. We investigated the following characteristics of M. sativa in natural and managed populations: floral traits, pollinator activity, and breeding system. Our results indicated the management could affect the number of flowers produced; however, there was no detectable effect on the seed set per flower. We found the percentage of seeds among pollinated flowers in the managed population was significantly higher than that in the natural population. Moreover, the increase in the proportion of pollinated flowers could significantly increase seed set per flower, and pollinator visitation rate was the important limiting factor for seed set in both populations. Andrena lebedevi Popov was found to be the most frequent pollinator in both populations. Outcrossing was dominant in the breeding system and insect pollination played an important role in outcrossing. Our study suggested that proper management (artificial selection) could promote pollination success of M. sativa.